1995] JOHNSON ЕТ AL.: BRUCНIDS AND PALM SEEDS
Principes, 39( \), \995, рр. 25-35 Bruchid Beetles and Palm Seeds: Recorded Relationships
1 2 1 3 CLARENC" Т)<'! JOHNSON, Scoтr Z(1NA, ANU JлN Л N!LSSON • 1Department of Biological Sciences, Northern Arizona University, Flagstaff, AZ 86011, USA 'Fairchild Tropical Garden, 11935 Old Cutler Road, Miami, FL 33156 and Department of Biological Sciences, Florida lnternational University, Miami, FL 33199, USA
ABSTRACT beetles. An examination of bruchids and their pa]m hosts is the suЬject of this paper. New World Bruchidae in the triЬe Pachymerini, suЬfamily Pachymerinae, alrnost exclusively feed in the seeds of palms А significant barrier to this undertaking is that and are therefore commonly referred to as palm bruchids. We most host records are. not substantiated Ьу seeds have found that these seed beetles have rnoderate preferences or voucher specimens of the palms from which for different taxa of palms. The bruchid ge:шs Caryobruchus the beetles were rea!'ed. Consequently, the accu shows definite preference for palms in the tribes Corypheae, Phoeniceae, and Hyophorbeae. Seeds of the Phytelephanto racy of the historical records cannot Ье verified. ideae and Сосоеае are fed upon Ьу the genera Caryoborus, Also, many of the names of the paJms have been Speciomerus, and especially species of Pachymerш. Thus changed since the original host records were pub palm bruchids do show specificity for host plants, and the lished or placed on insect labels. Many of the records reported in the literature are likely to Ье fairly accurate. puЬlished names of the paJm hosts are not those Among the many threats to the health and life in current use. of seeds, bruchid (seed) beetles (Coleoptera: Bru After assemЬ\ing the host records and correct chidae) are some of the most ubiquitous and for ing for the outdated bruchid and paJm taxonomy, midaЬle. These beetles lay their eggs on seeds or we found, simply Ьу examination, that there were fruits, and their larvae feed inside seeds, usually distinct relationships between genera of seed bee destroying the seeds in the process. From а plant's tles and some of the host genera of paJms. Thus, poiпt of view, а seed has only t\Vo purposes: to еvеп allowing for some spurious host records, 'Ne package the emЬryo for dispersal and to nurture have probaЬly estaЬlished that there are distinct the emЬryo until it сап establish itself as а seedling. host preferences between paJm bruchids and their From the bruchid's perspective, а seed is а nurs hosts. ·'"Y -curn-cafeteria, rich in food resources апd often \Vell protected from the environment. In short, it Plea for Bruchids Reared from is the perfect place for bruchid larvae to feed. Palm Seeds The conflict between these differing ageпda has W е strongly encourage paJm growers and spc led to some of the most interesting coevolutionary cialists to collect the insects that emerge froш the relationships in Ьiology. seeds of paJms and send them along \Vith а sample The fi.rst taxonomic revision of the bruchid tribe of the seeds to either CDJ, SZ, or JAN. Now that Pachymerini of the New World was recently pub the classification of paJm brucEds has been largely lished Ьу Nilsson and Johnson (l9Y3J. Included completed, we can identify the bruchids with а in their revision were published and unpuЬlished high degree of certainty. This is of value to paJm host records for these beetles, рrоЬаЫу all of which growers in controlling the insects. Also, now, more f. J on paJm seeds. With better understanding of than ever, ecological studies of these interesting i1ruchid taxonomy and relationships, \Ve are now plants and insects will Ье of greater value sincc in а position to examine the host records of these names can Ье applied to Ьoth plants and insects.
Bruchid Beetles 3 Present address: Central Arizona College, Apache Junction Caщpus, 273 Е. U.S. High''"Y 60, Apache Junction, AZ Beetles in the family Bruchidae feed in the seecls 85219, USA. of about 33 families of plants, but most feed in 26 PRINCIPES [VOL. 39 the Leguminosae. The reasons for host specificity About 84% of the hosts of bruchids are in the to any family are not easily ascertained, but some Leguminosae (J ohnson 197 О). Most of the other possiЬle reasons are discussed below. hosts are in the families Convolvulaceae (4.5%), Bruchids are in the superfamily Chrysomelo Arecaceae (4.5%), and Malvaceae (2%). Тhе other idea, а superfamily that includes the leaf beetles 5% are in 29 other families. Of the 5%, the and the longhomed wood borers. The larvae of ComЬretaceae, Rhamnaceae, Sterculiaceae, and the leaf beetles feed in or mine the leavcs, roots, especially the Tiliaceae host more bruchids than and stems. The larvae of the longhorned wood the other families. borers feed in wood and roots. According to John Seed beetles in the suЬfamily Pachymerinae are son ( 1981) it is likely that the chrysomeloid ances medium to large size seed predators · consisting tors of bruchids radiated onto the seeds of the chiefly of "primitive" forms. They appear to Ье Leguminosae (or possiЬly its ancestors) shortly after naturally distriЬuted in the tropics and subtropics it arose and then later evolved the ability to feed of all continents except Australia, and extend into in seeds of other families. All known seed beetles temperate zones in some areas. Palm bruchids are feed on, and usually live in, seeds. confined to the trihe Pachymerini, all of which are Тhе family Bruchidae consists of about 1,500 endemic to the Americas. Two species are now descriЬed species grouped in the suЬfamilies estaЬlished in Africa as tne result of human intro AmЬlycerinae, Bruchinae, Eubaptinae, К ytorhin ductions. Most species ~f Pachymerini are tropi inae, Pachymerinae, and Rhaebinae (Southgate cal, but species occur as far north as Texas and 1979). About 80% of bruchid species are in the as far south as Argentina. The Pachymerini almost Bruchinae, 10% in the AmЬlycerinae, 9% in the exclusively feed in palm seeds and are therefore Pachymerinae, with the other 1% assigned to the commonly called palm bruchids. Pachymerus other three suЬfamilies. abruptestriatus (Gyllenhal), however, has been · The life histories of bruchids vary consideraЬly reported to feed in the seeds of Diospyros sp. · in egg laying, larval entry, and larval feeding (Ebenaceae) (Bondar 1941). This record may Ье ·. behavior. The generallife cycle is that the adult spurious. The remaining two trihes of the Pachy· lays an egg on а fruit or seed, and the first stage merinae are native to the Old W orld and most (instar) larva bores through the fruit and/ or seed feed upon seeds of Leguminosae, some in the coat and enters the seed. The first stage larva is ComЬretaceae, with some questionaЫe reports from highly modified to enter seeds and has many spines the 1Jmbelliferae and Pandanaceae. and often well developed legs for this purpose. Once it has entered а seed it molts into а legless . Bruchid-Host lnteractions gruЬ and begins to feed. А larva usually molts Early coevolutionary studies between bruchids .· three more times as it continues to feed inside the and their hosts were conducted Ьу Janzen (1969) seed; usually only one, but sometimes several seeds who suggested traits that Leguminosae possess are eaten. Bruchids usually pupate inside the seed that seem to protect their seeds from seed beetles. that the larva has fed upon, but those tlpt feed Center and Johnson (197 4) suggested counter· on several seeds may build а pupal chamber Ьу mechanisms Ьу bruchids to these plant traits. gluing several seeds together, or some may leave Interactions between bruchids and toxic seeds have the seeds and spin а cocoon in which they pupate. been discussed Ьу several authors, but the research The larva starts and usually the adult finishes of Janzen (e.g., 1977, 1978) and Rosenthal (e.g., making а round exit hole in the seed and perhaps 1990) are especially enlightening. Ecological and the fruit through which the adult exits. Adult bru coevolutionary studies of palm bruchids and palmS chids feed on pollen and nectar and are not known known to us were made Ьу Janzen (1971), Wil;on to feed on seeds or fruits. Some species of bruchids and Janzen (1972), Smith (1975), Brown (1976), may complete many generations in а year, but Bradford and Smith (1977), Wright (1983), and most species of bruchids complete only one or а Smythe ( 1989). few generяtinns per year. ln tnflowers, pollen, and nectar JAN from specimens and the literature. SZ updat ~ .1 ---~-- =- '1' .. 1--1.". ry 'Тh.:. r1!1t~ in 'ГаЬlе 2 ·.. vere .. 1995] IOHNSON ЕТ AL.: BRUCHIDS AND PALM SEEDS 27
ТаЬlе 1. Bruchid beetles that have Ьееп reported to feed in seeds of Arecaceae. Palm taxa are grouped Ьу subfamily, tribe and genus. Cocos nucifera (coconut) is an unlikely host for palm \ruchids so record is unlikely and must Ье verijied. * = lntroduced into the New World. * * = Records are from hosts cultivated outside their native range.
Host Plant Palm Brucblds ARECACEAE. Coryphoideae Corypheae-31 genera; Cosmo. (26% of genera attacked) Thrinax Т. morrisii Н. Wendl. Caryobruchus gleditsiae Coccothrinax С. argentata (Jacq.) Bailey С. gleditsiae С. martii (Griseb. et Wendl.) Весе. С. gleditsiae Livistona *L. chinensis (Jacq.) R. Br. ех Mart. С. gleditsiae Serenoa S. repens (Bartram) Small С. gleditsiae Brahea 1 Brahea sp. С. rubidus С. veseyi В. armata S. Wats. С. veseyi В. brandegeei (Purpus) Н. Е. Moore С. veseyi Copernicia Copernicia sp. С. gleditsiae Pachymerus thoracicus С. alba Morong ех Morong & Britton Р. thoracicus С. hospita Mart. С. gleditsiae С. prunifera (Miller) Н. Е. Moore Р. bactris Р. sveni С. rigida Britton & Wilson ех Britton С. gledit'Siae Caryobruchus sp. С. sueroana Le6n С. gleditsiae С. tectorum (Kunth) Mart. Р. cardo С. macroglossa Wendl. еХ Весе. С. gleditsiae Washingtonia ** W. filifera (Linden) Wendl. С. gleditsiae * * W. robusta Wendl. С. gleditsiae c:abal Sabal sp. С. curvipes С. gleditsiae С. maya Speciomerus ruficornis S. bermudana L. Н. Bailey С. gleditsiae Caryobruchus sp. S. causiarum (0. F. Cook) Весе. С. gleditsiae Caryobruchus sp. S. domingensis Весе. С. gleditsiae Caryobruchus sp. S. etonia Swingle ех Nash С. gleditsiae Caryobruchus sp. maritima (Kunth) Burret С. gleditsiae Caryobruchus sp. S. mauritiiformis (Karst.) Griseb. & Н. Wendl. С. gleditsiae Caryobruchus sp. S. mexica.na Mart. С. curvipes С. gleditsiae Caryobruchus sp. Speciomerus ruficornis S. minor (Jacq.) Pers. С. gleditsiae Caтyobruchus sp. 28 PRINC!PES (VOL. 39
Table 1. Continued.
Host Plant Palm Bruchids
5. palmetto (Walt.) Lodd. ех J. А. & J. Н. Schult. С. gleditsiae С. marieae Caryobruchiis sp. 5. pumos (Kunth) Burret С. curvipes Caryobruchus sp. 5. rasei (0. F. Cook) Весе. С. gleditsiae С. veseyi Caryobruchus sp. S. uresaпa Trel. С. gleditsiae Caryobruchus sp. S. уара С. W right ех Весе. С. gleditsiae Caryobruchus sp. Phoeniceae-1 genus; African/ Asian (100%) Phoenix *Р. sylvestris (L.) Roxb. С. gleditsfae' Borasseae-7 genera; African/ AsiЭ.n (0%) Calamoideae Calameae-19 genera; African/ Asian (one species in the Americas) (0%) Lepidocaryeae-3 genera; American (33%) 1\llauritia М. jlexuosa L.f. Caryoborus gracilis Nypoideae-1 genus; Asian (0%) Ceroxyloideae Cyclospatheae-1 genus; American (О%) Ceroxyleae-5 genera; Cosmo. (0%) Hyophorbeae-5 genera; American (20%) Chamaedorea Chamaedorea sp. Caryobruchus gleditsiae С. elegans Mart. С. marieae С. maya С. seifГizii Burret С. gleditsiae Arecoideae Caryoteae- 3 genera; Asian (0%) lriarteeae-6 genera; American (17%) Dictyocaryum D. JUscum (Karst.) Н. Wendl. Caryoborus gracilis Podococceae-1 genus; African (О%) Areceae-86 genera; Cosmo. (5%) Euterpe Euterpe sp. Pachymerus bactris Е. oleracea Mart. Р. sveni Oenocarpus sp. Caryoborus gracilis О. Ьataua Mart. С. gracilis Areca. *А. triandra Roxburgh ех Buchanan-Hamilton Pachymerus cardo Сосоеае-22 genera; mostly American (3 African) (59%) Butia В. capitata (Mart.) Весе. Р. bridwelli Cocos С. nucifera L. Р. bactris Р. nucleorum 5уав.,J:и$ S. соrопыа (Mart.) Весе. .1!. ЫiCtHs Р. nucleorum 5. jlexuosa (Mart.) Весе. Р. bactris S. r·omrmzoffiana (Cham.) Glassman Р. bactris Р. cardo Р. nucleorum 1995] JOHNSON ЕТ AL.: BRUCHIDS AND PALM SEEDS 29
ТаЬlе 1. Continued.
Host Plant Palm Bruchids
S. schizophylla (Mart.) Glassman Р. bactris Р. nucleorum S. vagans (Bondar) Hawkes Р. bactris Р. nucleorum Attalea Attalea sp. Speciomerus giganteus S. rujicornis Р. bactris Р. cardo А. JU.nifera Mart. ех Sprengel Р. bactris Р. nucleorum А. lapidea (Gaerther) Burret Р. bactris Р. nucleorum А. spectabilis (Mart.) Burret Р. cardo А. tessmannii Burret Р. cardo А. victoriana Dugand Р. cardo Scheelea 1 S. gomphococca (Mart.) Burret Р. ca~do S. martiana Burret Speciomerus giganteus Pachymerus bactris S. brachyclada Burret Р. cardo S. excelsa Karst. S. giganteus S. leandroana Barb. Rodr. Pachymerus cardo S. liebmannii Весе. S. giganteus S. macrolepis Burret S. giganteus Pachymerш cardo S. maracaibensis (Mart.) Burret Р. cardo S. rostrata (Oersted) Burret S. giganteus Р. cardo S. zonensis L. Н. Bailey S. giganteus Р. cardo OrЬignya OrЬignya sp. Р. bactris Р. nucleorum О. cohune (Mart.) Dahlgren ех Standley Р. bactris Р. cardo Р. nucleorum О. phalerata Mart. Р. bactris Р. cardo Р. nucleorum S. giganteus Maximiliana Maximiliana sp. Caryoborus serripes М. maripa (Correa) Drude Р. cardo Elaeis *Е. guineensis Jacq. Р. bactris Е. oleifera (Kunth) Cortes Р. cardo Acrccomia Acrocomia sp. Speciomerus revoili Р. nucleorum А. aculeata (Jacq.) Lodd. S. revoili Р. bactris Р. cardo Р. nucleorum А. totai Mart. S. revoili Р. nut.:leorum Aiphanes А. aculeata Wilid. Р. cardo Bactris В. caryotifolia Mart. Р. sveni 30 PRINCIPES [VOL. 39
ТаЫе 1. Continued.
Host Plant Palm Bruchids
В. cuesa Crueger ех Griseb. Р. cardo В. maraja Mart. Р. sveni В. guineensis (L.) Н. Е. Moore S. ruficornis Р. bactris В. gasipaes Kunth Р. сагdо "" ,;., Desmoncus Desmoncus sp. Р. sveni D. polyacanthos Mart. Р. bactris Astrocaryum Astгocaryum sp. Caryoborus serripes S. ruficornis А. huicungo Dammer ех Burret С. serripes А. standleyanum L. Н. Bailey Р. bactris Geonomeae-6 genera; American (О%) Phytelephantoideae-3 genera; American (33%) Phytelephas Phytelephas sp. Caryobor~s chirique.,;,si-5 Р. aequatorialis Spruce С. chiriquensis Р. macrocarpa Ruiz & Pavon С. chiriquensis used to produce ТаЬ!е l. ТаЬ!е l shows host example, species of palms formerly placed in Cocos preferences even though puЬ!ished data were orig are now disposed in more than five different gen inally thought to Ье marginal in validity. These era. Thus, "Cocos sp." as а host record is rather data show relationships between classifications of uninformative. Other puzzling records are those both bruchids and palms. that use names with no botanical standing (viz., Some early host records are meaningless. F or nomina nuda, invalid comЬinations, typographical
1. Seeds of Copernicia sp. with Caryobruchus gleditsiae emerging from an exit hole in the seed on the left and а bruchid e:xit hole in the seed on the right. Note the typical round exit holes rnade Ьу these insects. Photo Ьу S. Zona. 1995] JOHNSON ЕТ AL.: BRUCHIDS AND PALM SEEDS 31
ТаЫе 2. Recorded hosts Jor New World species ofраlт bruchids (Pachyтeriпi). Most host records reported Ьу Nilssoп & ]ohnson (1993) were from seeds апd plants collected Ьу and identified Ьу Jthers. Because CDJ has used the host records of Zacher (1952) Jor studies in bruchids that Jeed in leguтes апd has Jouпd тапу of theт to Ье reliaЬle, we Jeel Zacher's records to Ье тате reliaЬle thaп тost others that have been puЬlished. We do not know, however, where he obtained тапу of his records. Тhе reports Ьу Udayagiri & Wadhi (1989) and other authors are mostly from the literature. lп .this. taЬle, the пате< (>J the. host plaпts are a.s шritten iп the ariginal puЬlications. For ease of reference to the literature we thiпk that the патеs reported iп the literature should Ье preseпted here еvеп though sоте are nomina nuda от invalid. Тherefore, we have placed valid патеs iп this taЬle Jollowed Ьу the invalid патеs iп parentheses Jollowed Ьу the literature citatioп iп which the iпvalid пате was reported. Nomina nuda are in quotes and not italicized. Names of раlт bruchids are in bold.
L.Jryoborus chiriquensis Sharp Host Plants.-Phytelephas sp.: Nilsson & Johnson 1993:16. Р. aequatorialis: Nilsson & Johnson 1993:16. Р. macrocarpa: Zacher 1952:470; Udayagiri & Wadhi 1989:239. Caryoborus gracilis Ni1sson / Host Plants.-Dictyocaryum fuscum: Nilsson & Johnson 1993:17. Oenocarpus sp.: NЦsso~ & Johnson 1993:17 (as ]essenia sp.: Ni1sson & Johnson 1993:16). О. bataua: (as ]essenia bataua: Ni1sson & Johnson 1993:17). Mauritia jlexuosa (as М. fiexulosa: Ni1sson & Johnson 1993:16). Palm Seeds: Nilsson & Johnson 1993:17. Caryoborus serripes (Sturm) Host Plants.-Astrocaryum sp.: Bridwell 1929:154; Zacher 1952:469; Udayagiri & Wadhi 1989:238; Nilsson & JohnscYJ 1993:20. А. huicungo: Nilsson & Johnson 1993:20. Maximiliana sp.: Zacher 1952:470. C··"!obruchus curvipes (Latreille) .юst Plants.-Cocos sp. (?): Bridwell 1918:493. Sabal sp.: Bridwell 1929:156; Zacher 1952:470; Zona 1990:614. Sabal sp. (as lnodes sp.: Bridwell 1929:156; Udayagiri & Wadhi 1989:239). 5. mexicana (as lnodes texana: Bridwell 1918:493; Zacher 1952:470; Udayagiri & Wadhi 1989:239). 5. pumos: Nilsson & Johnson 1993:23. Caryobruchus gleditsiae (Johansson and Linne) Host Plants.-Chamaedorea sp.: Nilsson & Johnson 1993:25. С. seifrizii: Nilsson & Johnson 1993:25. Coccothrinax argentata: Woodruff 1968:1; Nilsson & Johnson 1990:53; Nilsson & Johnson 1993:25. С. martii: Woodruff 1968:1. Copernicia sp.: Nilsson & Johnson 1990:53; Nilsson & John.son 1993:25. С. hospita: Nilsson & Johnson 1993:24. "Copernicia inderog1ossa": Nilsson & Johnson 1993:25. С. macroglossa (as "С. torram" and С. torreana: Nilsson & Johnson 1993:25). С. rigida: Nilsson & Johnson 1993:25. С. sueroana: Nilsson & Johnson 1993:25. Livistona chinensis: Woodruff 1968:1; Nilsson & Johnson 1990:53; Nilsson & Johnson 1993:25. Phoenix sylvestris: Woodruff 1968:1; Nilsson & Johnson 1990:53; Nilsson & Johnson 1993:25. Sabal sp.: Riley & Howard 1892:166; Cushman 1911: 504; Zacher 1952:470; Udayagiri & Wadhi 1989:239; Zona 1990 614; Nilsson & Johnson 1993:28. S. bermudana: Nilsson & Johnson 1990:53; Nilsson & Johnson 1993:24.5. causiarum: Nil. .. & Johnson 1990:53; Nilsson & Johnson 1993:25. 5. domingensis: Nilsson & Johnson 1990:53; Nilsson & Johnson 1993:25 (as lnodes neglecta: Nilsson & Johnson 1993:25). S. etonia: Nilsson & Johnson 1990:53; Nilsson & Johnson 1993: 25. S. maritima (as S. jlorida: Nilsson & Johnson 1993:25). S. mauritiiformis (as S. glaucescens: Woodruff 1968:1; as S. mauritiaeformis: Woodruff 1968: 1). S. minor (а• 5. glabra: Nilsson & Johnson 1990:53; Nilsson & Johnson 1993:26). "Sabal longipedunculata": Nilsson & Johnson 1990:53; Nilsson & Johnson 1993:25. S. mexicana: Nilsson & Johnson 1993:25 (as lnodes texana: Zacher 1952:468; Udayagiri & Wadhi 1989:239; as Sabal texana: Nilsson & Johnson 1990:53). S. minor: Paxson 1961:75; Woodruff 1968:1; Nilsson & Johnson 1990:53. S. palmetto: Woodruff 1968:1; Nilsson & Johnson 1990:53; Zona 1990:615; Nilsson & Johnson 1993:25 (as 5. parvijlora: Woodruff 1968: 1; Ni1sson & Johnson 1990:53; Nilsson & !ohnson 1993:25). S. rosei: Nilsson & Johnson 1990:53; Nilsson & Johnson 1993:25. S. uresana: Pfaffenherger & Johnson 1976:34; Nilsson & Johnson 1990:53; Zona 1990:615; Nilsson & Johnson 1993:25. S. уара: Woodruff 1968:1; Nilsson & Johnson 1990:53; Nilsson & Johnson 1993:25. Serenoa repens: Woodruff 1968:1. Thrinax morrisii (as Т. microcarpa: Nilsson & Johnson 1990:53; Nilsson & Johnson 1993:25). Washingtonia filifera: Nilsson & Johnson 1990:53. W. robusta: Nilsson & • !ohns· \993:26. Pahn Seeds: Nilsson & Johnson 1993:25. Cary -;-..,., uchus marieae Nilsson and Johnson ;,. Host Plants.-Chamaedorea elegans: Nilsson & Johnson 1993:28. Sabal sp.: Nilsson & Johnson 1990:55. S. palmetto (as S. parvi.ftora: Nilsson & Johnson 1990:55; Nilsson & Johnson 1993:28).
maya NiЬson Host Plants.-Chamaedorea elegan,; Nilsson & Johnson 1993:28. Sabal sp.: Nilsson & Johnson 1993:28. rubldus (Chevro1at) fiost Plants.-Brahea sp.: Nilsson & Johnson 1993:31. 32 PRINCIPES [VOL. 39
ТаЬlе 2. Continued.
Caryobruchus veseyi (Horn) Host Plants.-Brahea sp. (as Erythea sp.: Bridwell 1929:157). В. armata (as Erythea armata: Nilsson & Johnson 1993: 32). В. brandegeei (as Erythaea brandegeei: Zacher 1952:469 and Erythea brandegeei; Bridwell 1929:157; Udayagiri & Wadhi 1989:241; Nilsson & Johnson 1993:32). Sabal rosei: Nilsson & Johnson 1993:32. Caryobruchus sp. Host Plants.-Copernicia rigida: Zacher 1952:470. Sabal bermudana: Zona 1990:614. S. causiarum: Plant Quarantine Dec. 1932:194; Zona 1990:614. S. domingensis: Zona 1990:614. S. etonia: Zona 1990:614. S. maritima. Zona 1990:614. S. mauritiiformis: Zona 1990:614. S. mexicana: Zona 1990:614. S. minor: Zona 1990:614. S. palmetto: Zona 1990:614. S. pumos: Zona 1990:614. S. rosei: Zona 1990:614. S. uresana: Zona 1990:614. S. уара: Zona 1990:614. Speciomerus giganteus (Chevrolat) Host Plants.-Attalea sp.: Bridwell 1929:150; Zacher 1952:469; Udayagiri & Wadhi 1989:240; Nilsson & )ohnson 1993: 36. OrЬignya phalerata (as О. martiana: Zacher 1952:469; Udayagiri & Wadhi 1989:240; as Scheelea phalerata: Silva 1977:2; as "Вahassu nut": Nilsson & Johnson 1993:36). Scheelea excelsa: Bridwell 1929:150; Zacher 1952:470; Udayagiri & Wadhi 1989:241. S. liebmannii: Nilsson & Johnson 1993:36. S. macrolepis: Nilsson & Johnson 1993:36. "Scheelea orhignya": Zacher 1952:469; Udayagiri & Wadhi 1989:240. S. martiana (as Attalea excelsa: Zacher 1952:469). S. rostrata; Janzen 1971 :93; Janzen 1980:947; Udayagiri & Wadhi 1989:239; Nilsson & Johnson 1993:36. r· zonensis: Zacher 1952: 470; Udayagiri & Wadhi 1989:239; Nilsson & Johnson 1993:36. Speciomerus revoill (Pic) Host Plants.-Acrocomia sp.: Bridwell 1929:150; Zacher 1952:469; Udayagiri & Wadhi 1989:238. А. aculeata (as А. sclerocarpa: Silva 1977 :2). А. totai: Nilsson & Johnson 1993:38. Speciomerus ruficornis (Germar) Host Plants.-Astrocaryum: Bridwell1929:151; Zacher 1952:469; Udayagiri & Wadhi 1989:240. Attalea sp.: Silva 3, Bactris guineensis (as В. minor: Silva 1977:3; Udayagiri & Wadhi 1989:241). Cocos sp.(?): Bridwell 1918:493. Sabal (as Inodes sp.: Bridwell 1929: 156; Udayagiri & Wadhi 1989:239). Sabal sp.: Bridwell1929:156; Zacher 1952:470. S. mexicana.: (as lnodes texana: Bridwell 1929:156; Zacher 1952:470; Udayagiri & Wadhi 1989:239). Pachymerus abruptestriatus (Gyllenhal) Host Plants,-Diospyros sp.: Bondar 1941:303; Zacher 1952:469; Prevett 1966Ь;187; Udayagiri & Wadhi 1989:242; Nilsson & Johnson 1993:45.
Pachymerцs bactris (Linne) Host Plants.-Acrocomia aculeata (as А. sclerocarpa: Bridwell 1929:160; Zacher 1952:469; Silva 1977:4; Udayagiri & Wadhi 1989:244). Astrocaryum standleyanum: Nilsson & Johnson 1993:47. Attalea sp.: Bridwell 1929:160. А. funifera: Zacher 1952:469; Udayagiri & Wadhi 1989:245. А. lapidea (as Cocos lapidea: Zacher 1952:469; Udayagiri & Wadhi 1989: 245). Bactris guineensis (as В. minor: Zacher 1952:469; Udayagiri & Wadhi ~ 989:243; Nilsson & )ohnson 1993:47). Cocos nucifera: Zacher 1952:469; Udayagiri & Wadhi 1989:245. Copernicia prunifera (as С. cerifera: Bridwell1918:493; Hoffmann 1945:25; Zacher 1952:469; Udayagiri & Wadhi 1989:243; as Carnauba-Palme: Zacher 1952:470). Desmoncus polyacanthos: Zacher 1952:470; Udayagiri & Wadhi 1989:243. Elaeis guineensis: Zacher 1952:469; Silva 1977:3; Udayagiri & Wadhi' 1989:244, 245. Euterpe sp.: Zacher 1952:469; Udayagiri & Wadhi 1989:243. OrЬignya sp.: Zacher 1952:470. О. cohuм (as Attalea cohune: Zacher 1952:469; Udayagiri &; Wadhi 1989:245). О. phalerata (as О. martiana: Zacher 1952:470; Udayagiri & Wadhi 1989:245). О. speciosa (?, according to current palm classification, the valid name for this species is either · О. phalerata or О. cohune): Udayagiri & Wadhi 1989:244, 245. Scheelea martiana (as Attalea excelsa: Zacher 1952:469; Udayagiri & Wadhi 1989:244). Syagrus coronata (as Cocos coronata: Udayagiri & Wadhi 1989:245). S. jlexuosa (as Cocos · campestris: Zacher 1952:469; Udayagiri & Wadhi !989:243). S. romanzoffiana (as Cocos romanzoffiana: Zacher 1952:469; Udayagiri & Wadhi 1989:245). S. schizophylla (as "Licurioba campanemae": Zacher 1952:470; Udayagiri & Wadhi 1989: 245). S. vagans (as Cocos vagans Zacher 1952:4Ь9; Udayaglfi & Wadhi 1989:245). Pachymerus bridwelli (Prevett) Host Plants.-Butia capitata; Prevett 1966а:83; Nilsson & Johnson 1993:49; (as "Butiaca pitata" Udayagiri !989:238). Pachymerus cardo (Fiihraeus) Host Plants.-Acrocomia aculeata (as А. sclerocarpa; Nilsson & Johnson 1993:52). Aiphanes aculeata (as Moort
ТаЬlе 2. Continued.
melanococca: Nilsson & Johnson 1993:51). "'Englerophoenix sp.": Nilsson & Johnson 1993:52. Maximiliana maripa (as М. caribaea and М. regia Nilsson & Johnson 1993:52, 51). "Orhignya graciosa": Nilsson & Johnson 1993:51. OrЬignya phalerata (as Attalea speciosa: Nilsson & Johnson 1993:52). 5cheelea Ьrachyclada: Nilsson & Johnson 1993:52. 5. gomphococca (as Attalea gomphococca: Nilsson & Johnson 1993:52). 5. leandroana: Ni!sson & Johnson 1993:51. 5. macrolepis: Nilsson & Johnson 1993:52. 5. maracaibensis: Nilsson & Johnson 1993:52. 5. rostrata: Janzen 1971:93. 5. zonensis: Nilsson & Johnson 1993:52. 5yagrus romanzo.ffiana (as Cocos romanzo.ffiana: Nilsson & Johnson 1993:51). Pachymerus nucleorum (Yabricius) Host Plants.-Acrocomia sp.: Nilsson & Johnson 1993:54. А. aculeata (as А. sclerocarpa: Nilsson & Johnson 1993:54). А. totai: Nilsson & Johnson 1993:54. Attalea JUnifera: Zacher 1952:469; Udayagiri & Wadhi 1989:245. А. lapidea (as Cocos lapidea: Zacher 1952:469; Udayagiri & Wadhi 1989:245). Cocos nucifera: Zacher 1952:469; Udayagiri & Wadhi 1989: 245. Elaeis guineensis: Zacher !952:469; Silva 1977:3; Udayagiri & Wadhi 1989:245. OrЬignya sp.: Zacher 1952:470; Nilsson & Johnson 1993:54. О. cohune (as Attalea cohune: Zacher 1952:469; Udayagiri & Wadhi 1989:245). О. phalerata (as Attalea speciosa: Nilsson & Johnson 1993:54, OrЬignya martiana: Zacher 1952:470; Udayagiri & Wadhi 1989:245 and '1. speciosa: Zacher 1952:469; Udayagiri & Wadhi 1989:245; and as Babassu nuts: Nilsson & Johnson 1993:54). 5yagrus coronata (as Cocos coronata: Udayagiri & Wadhi 1989:245; Nilsson & Johnson 1993:54). S. romanzoffiana (as Cocos romanzo.ffiana: Zacher 1952:469; Udayagiri & Wadhi 1989:245). 5. vagans (as Cocos vagans: Zacher 1952:469; Udayagi1·i & Wadhi 1989:245). S. schizophylla (as "Licurioba campanemae": Zacher 1952:470; Udayagiri & Wadhi 1989:245). Pachymerus sveni Nilsson / Host Plants.-Bactris caryotifolia: Nilsson & Johnson 1993:55. В. maraja (as "В. masaja"; Nilsson & Johnson 1993:55). Copernicia prunifera (as С. cerifera: Nilsson & Johnson 1993:55). Desmoncu.s sp.: Nilsson & Johnson 1993:55. Euterpe oleracea: Nilsson & Johnson 1993:55.
Pachymerus thoracicus Prevett Host Plantso-Copernicia sp.: Nilsson & Johnson 1993:58. С. alba (as С. australis Zacher 1952:470; Prevett l966b:l90; Silva 1979:249; Nilsson & Johnson 1993:58). "С. orientalis": Prevett l966b: 190; Udayagiri & Wadhi 1989:245; Niliшor, & 0: o'шson 1993:58.
errors, etc.). These proЬ!ematic records are not Asian-Pacific, but four of the ten American genera included in ТаЬ\е l but are retained in ТаЬ\е 2 are fed upon Ьу bruchid beetles. to try to make sense of the names in the palm The bruchid genus CaryoЬruchus shows defi Ьruchid literature. nite preference for palms in the triЬes Corypheae, The percentage of genera per triЬe that are fed Phoeniceae (cultivated), and Hyophorbeae, The upon Ьу bruchids is in parentheses in ТаЬ\е l. typical bruchid extreme generalist is С. gleditsiae which has been reported to feed in seeds of nine Results and Discussion different genera, lnteractions Ьetween Palms and Palm Bru The seeds of Phytelephantoideae and especially chids. As the palm bruchids are American, it is Сосоеае are fed upon Ьу the genera CaryoЬorus, not unexpected that they have no host records Speciomerus, and especially species of Pachy from those suЬfamilies and tribes that are African merus. That these three genera of bruchids feed and/ or Asian (Borasseae, Ca\ameae, Nypoideae, in similar plants may not show any phylogenetic Caryoteae, and Podococceae), Species of Phoenix, affinities between bruchids and the phylogenetic an African-Asian genus and the only genus of the affi.nities of palms, Although а thorough phylo Phoeniceae, host bruchids when these palms are genetic analysis was not made of the palm bruchids cu]tivated in the Americas, It is curious that some Ьу Nilsson and Johnson (1993), CaryoЬorus and AIYc :rican triЬes have seldom or never been Speciomerus seem to Ье more close\y related to reported to host bruchids. Bruchids are not known CaryoЬruchus than to Pachymerus. to feed in the seeds of members of the С yclos The four groups most fed upon Ьу bruchids, patheae, Ceroxyleae, or Geonomeae. Host records the Сосоеае, Corypheae, Phytelephantoideae эдd from the Hyophorbeae, which includes the diverse о о Lepidocaryeae, have seeds that are quite difl:'erent and widespread Chamaedorea, are surprisingly in terms of seed morphology and chemistry. Most rare, The Areceae are predominantly African- palm seeds contain significant amounts of lipids 34 PRINCIPES
(triglycerides) and/or mannans (polysaccharides). JOHNSON, С. D. 1970. Biosystematics of the Arizona, Cal. Mannans, in one form or another, are character ifornia, and Oregon Species of the Seed Beetle Genus Acanthoscelides Schilsky (Coleoptera: Bruchidae). Univ. · istic ofpalm seeds (Daud and Jarvis 1992). Сосоеае Cslif. PuЬl. Entomol. 59: 1-116. have oil-rich endosperms and heavy endocarps. ---. 1981. Seed beetle host specificity and the system. Corypheae have thin endocarps and oil-poor endo atics of the Leguminosae, рр. 995-1027, + 61 рр. sperms. Phytelephantoideae have thick, woody mtcrofiche: In: R. М. Polhtll and Р. Н. Raven (eds.). pericarps, thin endocarps and oil,poor endo Advances rn Legume Systematics. Royal Botanic Garderis Kt:Jw. xvi + 105G рр. in two parts. ' sperms. Lepidocaryeae have fruits that are oily NILSSON, J. А. AN~ С. D. JoHNSON. 1990. А new species and juicy, the endocarp is thin, and the seed not of palm bruch1d from Cuba and а redescription of Саг. especially oil-rich. The epicarp is covered with yobruchus gleditsiae (L.) (Coleoptera: Bruchidae: Pachy. hard, shiny overlapping scales. Because species of merinae). Coleopts. Bull. 44(1): 50-59. --- AND . 1993. А taxonomic revision of the Pachymerus feed predominantly in Сосоеае (ТаЬ!е palm bruchids (Pachymerini) and а descripiion of the l ), it may Ье that most species are adapted to world genera of Pachyrnerinae (Coleoptera: Bruchidae: feeding in seeds rich in oil and with heavy endo Pachymerinae). Memoirs of the Arnerican Entomological carps. Apparently CaryoЬruchus, Speciomerus, Society. 41. 104 рр. and CaryoЬorus are mostly adapted to seeds with PAXSON, D. W. 1961. Letters. Principes 5: 75-76. little oil and with thin endocarps. PFAFFENBERGER, G. S. AND С. D, JOHNSON. 1976. Biosyste. matics of the first-stage lar1Vae of some North American Bruchidae (Coleoptera). U.S. Dept. Agric. Tech. Bull. LIТERATURE CIТED 1525. 75 рр. РRЕVЕТТ, Р. F. 19бба. А new genus and species of BONDAR, G. 1941. Notas entomo!Ogicas da Baia. VII. 4. merinae from South America. Proc. Entomol. Soc. Um Bruchideo novo do genero Pachymerus. Rev. Ento~ (В) 35: 81-83. mol. 12: 302-303. ---. 1966Ь. The identity of the palm kernel Ьorer BRADFORD, D. F. AND С. С. SмiTH. 1977. Seed predation Nigeria, with systematic notes on the genus Pach:ymerui and seed number in Scheelea palm fruits. Ecology 58(3): Thunberg (Coleoptera: Bruchidae). Bull. Entomol. 5 667-673. 181-192. BRIDWELL, J. С. 1918. Notes оп the Bruchidae and their ---. 196 7. N otes on the Ьiology, food plants and dis· parasites in the Hawaiian Islands. Proc. Hawaiian Ento tribution of Nigerian Bruchidae (Coleoptera), with partic mol. Soc. (1917)3: 465-505. ular reference to. the northern region. Бull. Entomol. Soc. ---. 1929. 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Bruchids and legumes: economics, 4(6): 489-510. ecology and coevolution. Proceedings of the Second lnter DAUD, М. J. AND М. С. )ARVIS. 1992. Mannan of'>il palm national Symposium on Bruchids and Legumes (ISBL-2) kernel. Phytochemistry 31: 463-464. held at Okayama (Japan), September 6-9, 1989. Кluwer HoFFMANN, А. 1945. Coleopteres bruchides et anthriЬides. Academic PuЬlishers, Dordrecht, Boston, London. 407 Faune de France 44:1-184. рр. JANZEN, D. Н. 1969. Seed-eaters versus seed size, number, SILVA, Р. 1977. Corre<;oes е aditamentos ао Quarto CatЗ.logo toxicity, and dispersal. Evolution 23: 1-27. dos insetos que vivem nas plantas do Brasil (Coleoptera, ---. 1971. The fate of Scheelea rostrata fruits beneath Bruchidae, Pachymerinae). Rev. Bras. Entomol. 21(1): the parent tree: predispersal attack Ьу bruch:ids. Principes 1-5. 15(3): 89-101. 1979. Pachymerus thoracicus Prevett (Coleop· ---. 1977. 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